Abstract [en]

1. The size of an individual is an important determinant of its trophic position and the type of interactions it engages in with other heterospecific and conspecific individuals. Consequently an individual's ecological role in a community changes with its body size over ontogeny, leading to that trophic interactions between individuals are a size-dependent and ontogenetically variable mixture of competition and predation.

2. Because differently sized individuals thus experience different biotic environments, invasion success may be determined by the body size of the invaders. Invasion outcome may also depend on the productivity of the system as productivity influences the biotic environment.

3. In a laboratory experiment with two poeciliid fishes the body size of the invading individuals and the daily amount of food supplied were manipulated.

4. Large invaders established persistent populations and drove the resident population to extinction in 10 out of 12 cases, while small invaders failed in 10 out of 12 trials. Stable coexistence was virtually absent. Invasion outcome was independent of productivity.

5. Further analyses suggest that small invaders experienced a competitive recruitment bottleneck imposed on them by the resident population. In contrast, large invaders preyed on the juveniles of the resident population. This predation allowed the large invaders to establish successfully by decreasing the resident population densities and thus breaking the bottleneck.

6. The results strongly suggest that the size distribution of invaders affects their ability to invade, an implication so far neglected in life-history omnivory systems. The findings are further in agreement with predictions of life-history omnivory theory, that size-structured interactions demote coexistence along a productivity gradient.

Schröder, Arne

Abstract [en]

Alternative stable states have been, based on theoretical findings, predicted to be common in ecological systems. Empirical data from a number of laboratory and natural studies strongly suggest that alternative stable states also occur in real populations, communities and ecosystems. Potential mechanisms involve population size-structure and food-dependent individual development. These features can lead to ontogenetic niche shifts, juvenile recruitment bottlenecks and emergent Allee effects; phenomena that establish destabilising positive feedbacks in a system and hence create alternative stable states.

I studied the consequences of population size-structure for community dynamics at different scales of system complexity. I performed laboratory and ecosystem experiments. Small poecilliid fishes and planktonic invertebrates with short generation times and life spans were used as model organisms. This allowed me to assess the long-term dynamics of the populations and communities investigated.

The main experimental results are: (a) An ontogenetic niche shift in individuals of the phantom midge Chaoborus made the population vulnerable to an indirect competitive recruitment bottleneck imposed by cladoceran mesozooplankton via rotifers. Consequentially the natural zooplankton food web exhibited two alternative attractors. (b) Body size determined the success of Poecilia reticulata invading resident population of Heterandria formosa and thus the type of alternative stable state that established. Small invaders were outcompeted by the residents, whereas large invaders excluded their competitor by predating on its recruits. (c) External juvenile and adult mortality altered the internal feedback structure that regulates a laboratory population of H. formosa in such a way that juvenile biomass increased with mortality. This biomass overcompensation in a prey population can establish alternative stable states with top-predators being either absent or present.

The major conclusion is that size-structure and individual growth can indeed lead to alternative stable states. The considerations of these ubiquitous features of populations offer hence new insights and deeper understanding of community dynamics. Alternative stable states can have tremendous consequences for human societies that utilise the ecological services provided by an ecological system. Understanding the effects of size-structure on alternative stability is thus crucial for sustainable exploitation or production of food resources.

Nilsson, Karin

Umeå University, Faculty of Science and Technology, Department of Ecology and Environmental Sciences.

2010 (English)Doctoral thesis, comprehensive summary (Other academic)

Abstract [en]

Most animals grow substantially during their lifetime and change in competitive ability, predatory capacity and their susceptibility to predation as they grow. This thesis addresses the implications of this on regulation and dynamics within populations as well as between population interactions.

In size-structured populations either reproduction or maturation may be more limiting. If juveniles are competitively superior, the competitive bottleneck will be in the adults and reproduction will be limiting. Mortality will in this case result in overcompensation in juvenile biomass through increased reproduction. Compensation in biomass was demonstrated in Daphnia pulex populations subjected to size-independent mortality, where juvenile biomass did not decrease when a substantial harvest was imposed due to increase per capita fecundity. This supported that juveniles were superior competitors and that population cycles seen in Daphnia are juvenile-driven.

Compensatory responses in biomass may lead to that predators facilitate eachothers existence by feeding on a common prey, a phenomenon coined emergent facilitation. In an experimental test of the mechanism behind emergent facilitation it was demonstrated that the invertebrate predator Bythotrephes longimanus was favoured by thinning of its prey Holopedium gibberum. The thinning mimicked fish predation and targeted large individuals while Bythotrephes preferrs small prey.

Size dependent predation also occurs within populations, i.e. cannibalism, were large individuals feed on smaller conspecifics. Two populations of the common guppy (Poecilia reticulata) originating from different environments were demonstrated to differ in cannibalistic degree. Cannibalism was also affected by the presence of refuges and females and juveniles from one population were better adapted to structural complexity than the other.

The effects of these differences in cannibalism on population regulation and dynamics were studied in long term population experiments. Both populations were regulated by cannibalism in the absence of refuges, and displayed cannibal-driven cycles with suppression of recruitment and high population variability. The presence of refuges decreased density dependence and population variability and harvesting of large females in the absence of refuges led to population extinctions in the more cannibalistic population. The less cannibalistic population had higher population biomass and stronger density-dependence in the presence of refuges. When refuges were present, cohort competition increased and cycles with short periodicity were seen.

Large individuals were not only cannibals, but could successfully prey on other species. Small and large guppies were allowed to invade resident populations of Heterandria formosa. Small invaders failed while large invaders succeeded as predation from large invaders broke up the competitive bottleneck that the resident population imposed on juveniles of the invader.